Zootaxa, Ephemeroptera and Plecoptera Biodiversity in Central

Ephemeroptera and Plecoptera biodiversity in central Patagonia, Chubut province, Argentina

PABLO PESSACQ1 & MARÍA LAURA MISERENDINO2 Laboratorio de Investigación en Ecología y Sistemática Animal (LIESA), Universidad Nacional de la Patagonia San Juan Bosco. Sarmiento 849, 9200. Esquel, Chubut, Argentina.E-mail: [email protected], [email protected]

Abstract

Aquatic insects in Central Patagonia (Andean region), Argentina, have received poor and fragmentary attention. In this paper we present the first preliminary inventory of Plecoptera and Ephemeroptera taxa for the area and include seasonal records of adult stages. We incorporate a set of environmental features for the sampling sites and biotopes as well as the main characteristics of the phytogeographical and biogeographical areas in which the species were found. Twelve species and eight genera of mayflies and eleven species and five genera of stoneflies are recorded for the first time in Chubut province. The stoneflies Neofulla biloba (Aubert), Alfonsoperla flinti McLellan & Zwick, Austronemoura eudoxiae Froehlich, Chilenoperla semitincta Illies, Neonemura illiesi Zwick and Potamoperla testacea Vera, and the mayflies Andesiops ardua (Lugo-Ortiz & McCafferty), Chaquihua bullocki Navás. And the family Oniscigastridae are new records for Argentina. The genera Americabaetis Kluge, Camelobaetidius Demoulin and Traverella Edmunds are confirmed for first time in the Andean region (Central Patagonia province) and four Plecoptera and two Ephemeroptera species constitute new records for the Patagonian steppe. We also include two new records for Ephemeroptera and Plecoptera for the Santa Cruz province. We reinforce the idea that patagonian aquatic environments have a highly spe- cialized and complex insect fauna which can be correlated with biogeographical aspects but also with a great environmental heterogeneity.

Key words: new records, inventory, Patagonia, Ephemeroptera, Plecoptera

Introduction

Knowledge of Ephemeroptera and Plecoptera fauna in Argentina is still poor and fragmentary (Domínguez 1998, Muzón & Bachmann 1998). This is certainly true for the Central and the Subandean Patagonia (Patago- nian subregion) in Chubut province, these areas had received little sampling effort. As a result of different research projects of the Laboratorio de Investigación en Ecología y Sistemática Animal (LIESA) several lotic environments in the area have been the object of intense and systematic collections (Miserendino 1996, 2001, Miserendino & Pizzolón 2000, 2004, Hollmann & Miserendino 2006). Even so, wide areas in the southern region, the Valdivian forest, and most of the steppe remain almost unexplored. At present, 25 mayfly species within 13 genera have been recorded for the Argentinean Subantarctic and Patagonian subregions (Andean region) (Domínguez et al. 1994, Muzón & Pessacq 2001, Nieto 2004). Six- teen of those species (in 8 genera) have been recorded in the Chubut province (Domínguez et al. 1994, Miser- endino & Pizzolón 2000, Miserendino 2001, Nieto 2004). Stoneflies are represented by 36 species within 27 genera (McLellan et al. 1990, 2006, Muzón & Bach- mann 1998, Nelson 1973), of which 17 (within 15 genera) have been recorded in the Chubut province (Aubert 1960, Illies 1963, McLellan et al. 2006, Miserendino 1996, Miserendino & Pizzolón 2000).

Accepted by R. DeWalt: 14 May 2008; published: 7 Jul. 2008 27

Material and methods

The present contribution is the result of several surveys carried out in 2005, 2006 and 2007 at 41 sampling sites (Table 1). The study area included the Middle West of Chubut province, the Chubut River basin, and the Santa Cruz River in Santa Cruz province, located within the Patagonian and Subantarctic subregions of the Andean region (Morrone 2006) (Fig. 1).

FIGURE 1. Sampling sites in Chubut province. Code of sampling sites in Table 1. P.N.: Los Alerces National Park.

For larval collections Surber, kick-net, and D-frame nets were used, whereas for adults aerial nets, pitfall traps, light traps and Malaise traps were employed (Hauer & Resh 1996). An environmental characterization for most of the sampling sites was performed. For that purpose, we compiled information previously published (Miserendino 2001, 2006, Miserendino & Pizzolón 2004) and new data. Substrate composition was estimated as percentage of boulder, cobble, gravel, pebble, and sand using a 1-m2 grid (Gordon et al. 1994). Stream order was obtained from Coronato & Del Valle (1988). Surface current speed was obtained by timing a bobber (average of 3 times) as it moved over a distance of 10 meters (Gordon et al. 1994). At each site we measured wet width and bank full width. The phytogeographical and biogeographical description was based on León et al. (1998) and Morrone (2006) respectively. The species composition of the riparian corridor was analyzed and main native and exotic species recorded.

Results

Environmental Characterization: The main environmental features of the sampling sites are presented in Table 2. This gives a picture of the diversity of watercourses visited during the study. Elevation of sampling sites ranged between 3 and 820 m. Stream order was between 2 and 6. Conductivity values ranged between 9 and 306 uS.cm-1. As expected, mountain streams showed lower conductivity values than those in the steppe. Sub-

strate size was similar across most mountain rivers, being mainly comprised of boulders and cobbles. Sub- strate size in steppe streams was finer, consisting mostly of cobbles, gravel, and sand.

TABLE 1. Sampling locations for Chubut and Santa Cruz provinces, Argentina. Sites 1-6, 8-19 and 26-30 were sampled on several occasions between May 2005 and March 2006, sites 20-25 were visited between October 2006 and April 2007. RN: Ruta Nacional, RP: Ruta Provincial.

Chubut Province Site Water Body Locality Other descriptor Lat. Long. Dates 1 Golondrinas Str. El Hoyo Intersection RP 258 41°59’33’’ 71°33’25’’ 2 Nameless Str. Puerto Patriada trib. Epuyen L. 42°08’17’’ 71°31’56’’ 3 Las Minas Str. Cholila Pre-urban reach 42°31’03’’ 71°25’13’’ 4 Las Minas Str. Cholila Urban reach 42°30’32’’ 71°25’34’’ 5 Esquel Str. Nr. Esquel Road access to “La Hoya’’ 42°50’28’’ 71°15’55’’ 6 Esquel Str. Nr. Esquel Pre-urban reach 42°53’48’’ 71°16’03’’ 7 Percey R. Alto Río Percey 15 km from Esquel 45°51’17’’ 71°25’23’’ 4-26/III/07 8 Percey R. Nr. Trevelin 2 km upstream Trevelin 43°04’22’’ 71°28’14’’ 9 Blanco Str. Nr. Los Cipreses At RN 259 intersection 43°10’38’’ 71°36’30’’ 10 Baggilt Str. Nr. Los Cipreses At RN 259 intersection 11 Nameless Str. Nr. Trevelin RP 17, Percey R. trib. 43°12’02’’ 71°24’01’’ 12 Nant y Fall Str. Nr. Trevelin At RP 17 intersection 41°13’24’’ 71°25’17’’ 13 Nameless Str. Nr. Los Ñires farm RP 17, Río Frío trib. 43°1’33’’ 71°26’36’’ 14 Río Frío Los Ñires farm Farm on SR 17 43°29’56’’ 71°20’22’’ 15 Nameless Str. Nr. los Ñires farm RP 17, Río Frío trib. 43°21’43’’ 71°30’22’’ 16 Glyn Str. Nr. Cerro Centinela Nearby RP 17 43°27’34’’ 71°33’25’’ 17 Carbón Str. Nr. Corcovado Pre-urban reach 43°32’11’’ 71°28’21’’ 18 Cabeza de Vaca Str. Nr. Corcovado At RP 17 intersection 43°29’58’’ 71°20’51’’ 19 Carbón Str. Corcovado Urban reach 43°32’07’’ 71°28’11’’ 20 Corcovado R. Nr. Corcovado El Palenque farm, RP 44 21 Poncho Moro Str. Nr. Poncho Moro farm RP 44 intersection 22 El Comisario Str. RP 44 intersection 43°44’45’’ 71°23’32’’ 23 Nameless Str. RP 44 & Corcovado trib. 43°44’36’’ 71°23’35’’ 24 Corcovado R. Oulet of Vintter L. 43°53’54’’ 71°25’28’’ 27/IV/07 25 Chubut R. Fofo Cahuel At RP 4 42°26’07’’ 70°30’35’’ 26 Chubut R. Piedra Parada At RP 12 42°40’25’’ 70°0.5’1’’ 27 Chubut R. Paso Berwin At RN 25 43°37’15’’ 68°57’05’’ 28 Chubut R. Las Plumas At RN 25 intersection 43°43’12’’ 67°17’21’’ 29 Chubut R. Valle Los Altares At RN 25 43°53’29’’ 68°23’5’’ 30 Chubut R. 28 de Julio Dwnstr. Ameghino dam 43°22’01’’ 65°53’17’’ 31 Coronado Str. Los Alerces Nat'l Pk. West side Rivadavia L. 42°37’60’’ 71°41’55’’ 12/III/07 32 Colegual Str. Los Alerces Nat'l Pk. Confluence Rivadavia R. 42°40’27’’ 71°41’55’’ 26/XI/06 33 Nameless Str. Los Alerces Nat'l Pk. N of Centinela str, RN 71 42°29’06’’ 71°23’21’’ 29/III/07 34 Nameless Str. Los Alerces Nat'l Pk. Pucon Pai camping., RN 71 42°29’44’’ 71°23’21’’ 29/III/07

35 Irigoyen Str. Los Alerces Nat'l Pk. Irigoyen Fall, RN 71 42°30’57’’ 71°21’43’’ 29/III/07 36 Nilson R. Los tres leones farm 2 km from Pico R. 44°15’06’’ 71°40’37’’ 29/I/07 37 Nameless str. Nr. Río Pico Access road to L. 2 28/IV/07 Santa Cruz Province 38 Santa Cruz R. Charles Für At RP 9 50°15’35’’ 71°53’40’’ V/05 39 Santa Cruz R. Condor Cliff At RP 9 50°13’07’’ 70°55’52’’ V/05 40 Santa Cruz R. Barrancosa At RP 9 50°30’30’’ 70°10’41’’ V/05 41 Santa Cruz R. Cte Luis Piedrabuena At RP 63 50°0’48’’ 69°0’32’’ V/05

TABLE 2. Environmental features of the sampling sites. Phytogeografical units codes: 1) Subantarctic province. 2) Pat- agonic province: Subandean district: a) Herbaceous steppe, occidental district: b) Herbaceus/shrub like steppe, central district c) Barren steppe d) Shrub steppe of Chuquiraga Avellanedae e) Shrub steppe of Junellia tridents. Ach: Austroce- drus chilensis, Nd: Nothofagus dombeyi, Na: N. antarctica, Np: N. pumilio, Mb: Maytenus boaria, S: Salix spp., Pn: Pop- ulus nigra, Pa: P. alba, Pm: Pseudotsuga mentziesii, Pc: Pinus contorta. Boul: boulder, Co: cobble, Pe: pebble, Gra: gravel, sand.

Site Stream Bank Width Substrate Current m/ Conductivity Elevation m Phytographic Riparian Veg. order (m) sec uS/sec Units 1 2 4.2 Co/Sand 2.6 126 361 1 Pm, Pc 2 1 1.8 Gra/Sand 3 76 495 1 Pm 3 2 3.25 Co/Boul 0.93 164 573 1 S 429Pe/Co3725551S 5 3 10.8 Boul/Co 3 105 721 2a Mb, Na 6 4 9 Co/Pe 0.9 184 610 2a S, Na 7 5 80 Boul/Co 1 80 600 1 S 8 5 100 Boul/Co 1.11 83 360 2a S 9 2 35 Boul/Co 0.4 16.8 450 1 Na 10 3 20 Boul/Co 0.6 21.6 460 1 Np, Sal 11 2 0.9 Gra/Sand 0.1 84 730 1 Na 12 3 31.5 Gra/Sand 1.45 113 690 1 Na 13 2 3.5 Co/Pe 3 9 670 1 Na 14 4 22.5 Sand/Gra 0.9 98 670 1 - 15 3 14.2 Boul/Co 1.5 87 700 1 - 16 2 5.66 Co/Pe 0.1 69 615 1 Na 17 3 10.5 Boul/Co 1.73 93 380 1 Pn, Pa 18 2 6.88 Co/Gra 1.06 63 700 1 Na, S 19 3 23.5 Boul/Co 1.35 84 400 1 S 20 4 50 Boul/co 1 - 430 1 S 21 3 12 Boul/Co 1 23.7 1 Na, Ach 22 2 4 Boul/Co 1.03 26 820 1 Na, Np 23 3 9 Boul/Co 0.8 29 770 1 Na, Np 24 4 30 Boul/Co 1.9 - 928 1 Na, Np 25 4 80 Pe/Gra 1.1-2.1 99 500 2a S 26 5 200 Co/Pe 0.9-1.2 148.7 440 2b S 27 5 160 Co/Sand 0.5-1.6 185.2 308 2d S

28 6 200 Pe/Gra/Sand 0.8-1.2 187.2 243 2d S 29 6 300 Boul/Co 1.7-1.9 189.5 300 2c S 30 6 70 Co/Pe 0.7-0.9 196 70 2c S, Pn, Pa 31 3 15 Co/Gra 0.8 - 541 1 Nd, Ach, S 32 3 25 Boul/Co 1.6 26.1 527 1 Nd, Ach 33 3 25 Boul/Co 0.7 41.2 580 1 Nd, Ach 34 3 3 Boul/Co 1.1 22.3 570 1 Nd, Ach, Ch 35 3 7 Boul/Co 1.3 26.1 570 1 Nd, Ach, Ch 36 3 10 Boul/Co 1.4 - 503 1 Nd, S 3737Gra/Co0.2- - 1 Nd 38 6 150 Co/Pe 0.35 - 170 2e - 39 6 200 Boul/Co 1.2 - 120 2e - 40 6 230 Pe/Gra 0.5 - 82 2e - 41 6 140 Pe/Gra 1.43 - 3 2e -

Ephemeroptera and Plecoptera analysis: This contribution increases to 24 the number of mayfly species that are known for the Chubut Province (within 15 genera), including the first records for Argentina of Andesiops ardua (Lugo-Ortiz & McCafferty 1999), Chaquihua Demoulin, 1955b and the family Onis- cigastridae. Moreover, the genera Camelobaetidius Demoulin, 1966, Americabaetis Kluge, 1992 and Traver- ella Edmunds, 1948 are recorded for the first time from the entire Andean Region (Table 3).

TABLE 3. Ephemeroptera and Plecoptera species recorded for Chubut province (including data from Santa Cruz River, Santa Cruz province). Site of collection, comments, previous records, and main literature are indicated when the taxa represent a new record. CH: Chubut province, NQ: Neuquén province, RN: Río Negro province, SC: Santa Cruz province. Records of Chaquihua & Siphlonella are in agreement with Domínguez et al. (2001) who mention the genus for southern Argentina in their generic key without further information.

Sites Comments Previous records References EPHEMEROPTERA Ameletopsidae Chaquihua bullocki (Navás 1, 2, 16 New generic record, Subant. subreg., S Chile Mercado & Elliot 2004 1930) Argentina Chiloporter eatoni Lestage 1931 Domínguez 1998 Baetidae Americabaetis alphus Lugo- 25, 26, 29 New generic record, S Am. trans. zone, Lugo-Ortiz & Mc Cafferty 1996 Ortiz & McCafferty 1996 Andean reg. Neotrop. reg., Argentina Andesiops ardua (Lugo-Ortiz & 29 New record, Argen- Subant. subreg., S Chile Lugo-Ortiz & Mc Cafferty 1999, McCafferty 1999) tina Nieto 2004 A. torrens (Lugo-Ortiz & Nieto 2004 McCafferty 1999) Camelobaetidius penai (Traver 25, 26, 29 New generic record, S Am. trans. zone reg., Domínguez 1998 & Edmunds 1968) Andean reg. Neotrop. reg., Argentina Caenidae Caenis sp. 3, 4, 5, 7, New family record, Neotrop. reg. Domínguez et al 1994, Miseren- 25, 26, 28 Andean reg. dino & Pizzolón 2000 Leptophlebiidae

Archethraulodes spatulus Pesca- 21 New generic record, Subant.& Patag. subreg., Domínguez 1998 dor & Peters 1982 Chubut prov. S Chile & Argentina (NQ) Dactylophlebia carnulenta 18, 20-22, New generic record, Subant. & Patag. sub- Domínguez 1998 Pescador & Peters 1980 37 Chubut prov. regs., S Chile & Argen- tina (NQ) Meridialaris chiloeensis 38, 40, 41 New record, Patago- Subant. & Patag. sub- Domínguez 1998 (Demoulin 1955a) nia steppe & Santa reg.s, S Chile & Argen- Cruz prov. tina (CH, NQ, RN) M. diguillina (Demoulin 1955c) Domínguez 1998 M. laminata (Ulmer 1920) 5, 6, 7, 8, 1st confirmed Subant.& Patag. subreg., Domínguez 1998, Miserendino 11,12,13,1 record, Chubut prov S Chile & Argentina (NQ, & Pizzolón 2000 4 RN) Nousia bella Pescador & Peters 9, 10, 12, 1st confirmed Subant. & Patag. subreg., Domínguez 1998, Miserendino 1985 20, 21, 37 record, Chubut prov S Chile & Argentina (NQ, 2001 RN) N. crena Pescador & Peters 1985 Domínguez 1998 N. delicata Navás 1918b Domínguez 1998 N. minor (Demoulin 1955c) Domínguez 1998 Penaphlebia vinosa (Demoulin 1 1st confirmed Patag. subreg., Argentina Domínguez 1998, Miserendino 1955c) record, Chubut prov. (NQ, RN) 2001 P. chilensis (Eaton 1883) 26 New record, Patago- Subant. & Patag. sub- Domínguez et al 1994 nia steppe regs., S Chile & Argen- tina (CH, NQ, RN) P. flavidula Pescador & Peters Domínguez 1998 1991 P. fulvipes (Needham & Murphy Domínguez 1998 1924) Rhigotopus andinensis Pescador Domínguez 1998 & Peters 1982 Traverella sp. 25-30 New generic record, S Am. trans. zone reg., Domínguez 1998 Andean reg. Neotrop. reg., Argentina Nesameletidae Metamonius anceps (Eaton Domínguez 1998 1885) Oniscigastridae Siphlonella sp. 7, 15, 16, New family record, Subant. subreg., S Chile Needham & Murphy 1924, Pictet 32 Argentina 1894 PLECOPTERA Eustheniidae Neuroperla schedingi (Navás Illies 1960b, Miserendino 1996 1929) Diamphipnoidae Diamphipnopsis samali Illies 1, 2 New generic record, Subant. & Patag. sub- Illies 1960b, Wais et al 1987 1960b Chubut prov. regs., S Chile & Argen- tina (NQ, RN) Austroperlidae Klapopteryx kuscheli Illies 38-41 New record, Patago- Subant. & Patag. sub- Illies 1960a, McLellan 2001 1960a nia steppe & Santa regs., S Chile & Argen- Cruz prov. tina (CH, NQ, RN)

Table 3. Continued Sites Comments Previous records References Penturoperla barbata Illies Illies 1960a, Miserendino 1996 1960a Griptopterygidae Alfonsoperla flinti McLellan & 15, 22,23 New generic record, Subant. subreg., S Chile McLellan & Zwick 2007 Zwick 2007 Argentina Antarctoperla michaelseni 28, 30, 38- New record, Patago- Subant. & Patag. sub- Illies 1963, Miserendino 2001 (Klapálec 1904) 41 nia steppe regs., S Chile & Argen- tina (CH, NQ, RN, SC) Aubertoperla illiesi (Froehlich Illies 1963 1960) Araucanioperla bullocki (Navás Illies 1963 1933) Ceratoperla fazi (Navás 1934) 22, 23 New record, Chubut Subant. & Patag. sub- Illies 1963 prov regs., S Chile & Argen- tina (RN) Chilenoperla semitincta Illies 5 1st confirmed Subant. subreg., S Chile Illies 1963, Miserendino 2001, 1963 record, Argentina Miserendino & Pizzolón 2000 Limnoperla jaffueli (Navás 29, 38-41 New record, Patago- Subant. & Patag. sub- Illies 1963, Miserendino 2001 1928) nia steppe regs., S Chile & Argen- tina (CH, NQ, RN) Notoperla archiplatae (Illies Illies 1958 1958) N. fasciata McLellan 2006 McLellan 2006 N. magnaspina McLellan 2006 McLellan 2006 Notoperlopsis femina Illies 1963 26 New record, Patago- Subant. & Patag. sub- Illies 1963, McLellan et al 2006 nia steppe regs., S Chile & Argen- tina (CH) Pelurgoperla personata Illies Illies 1963 1963 Potamoperla myrmidon (Mabille Illies 1963 1891) P. t est ace a Vera 2006 22, 23 New record, Argen- Subant. subreg., S Chile Vera 2006 tina Rhithroperla rossi (Froehlich Froehlich 1960, Illies 1963 1960) Senzilloides panguipullii (Navás Illies 1963 1928) Notonemouridae Austronemoura chilena Aubert 15, 21 22, New generic record, Subant. subreg., S Chile Aubert 1960, Illies 1961 1960 33-35 Chubut prov & Patag. subreg., Argen- tina (RN) A. eudoxiae Froehlich 1960 15 New record, Argen- Subant. subreg., S Chile Froehlich 1960, Illies 1961 tina Neofulla biloba (Aubert 1960) 35 New generic record, Subant. subreg., S Chile Aubert 1960, Illies 1961 Argentina Neonemura barrosi Navás 1919 21, 31 New generic record, Subant. subreg., S Chile Illies 1961, Wais et al 1987 Chubut prov & Patag. subreg., Argen- tina (RN, NQ) N. illiesi Zwick 1972 10 New record, Argen- Subant. subreg., S Chile Zwick 1972 tina

Udamocercia frantzi Illies 1961 5, 21, 22, New generic record, Subant. subreg., S Chile Illies 1961, Wais et all 1987 24, 37 Chubut prov & Patag. subreg., Argen- tina (NQ, RN) Perlidae Kempnyella genualis (Navás Illies 1964 1918b) Pictetoperla gayi (Pictet 1841) Illies 1964

The number of stonefly species now known from Chubut Province stands at 28 (in 23 genera). Moreover, The genus Neofulla Claassen, 1936, Alfonsoperla flinti McLellan & Zwick, 2007, Austronemoura eudoxiae Froehlich, 1960, Neonemura illiesi Zwick, 1972, Chilenoperla semitincta Illies, 1963 and Potamperla testacea Vera, 2006 are recorded for the first time in Argentina. Additionally, four species of stoneflies and two species of mayflies are recorded for the first time in Pat- agonia steppe. For Santa Cruz Province a new Ephemeroptera specific record and a new Plecoptera generic record are included. Table 3 provides a summary of the Ephemeroptera and Plecoptera inventory for Chubut province, including site of collection, comments, previous records and main references for every new record.

Discussion

The emergence periods of Ephemeroptera and Plecoptera species are predictable and well known in the northern hemisphere and in some southern hemisphere regions (Hynes & Hynes 1975, Scrimgeour 1991, Richard- son 2001). Unfortunately, literature about emergence periods of mayflies and stoneflies in Patagonia is limited to that of Miserendino & Hollmann (2005) and Hollmann & Miserendino (2008). The emergence data presented in this paper are not the result of a systematic sampling, nevertheless, the capture dates of adults constitute new information for the species in the area (Table 4). Mayflies emerged nearly exclusively in southern hemisphere in spring and summer, with the minor exception of Nousia delicata Navás, 1918a, which was also present in early autumn. Meridialaris chiloeensis (Demoulin 1955a) was the first mayfly to emerge, beginning in early October, which is consistent with reports by Hollmann & Miserendino (2006) who observed a bivolt- ine pattern with spring and summer generations for this species. Some stoneflies began emergence in late winter, but the majority waited until spring, with several emerg- ing throughout the summer months. Adults of the three Notonemouridae genera Austronemoura Aubert, 1960, Neofulla and Udamocercia Enderlein, 1909 were found mainly by the end of summer and beginning of autumn. Most Gripopterygidae emerged during spring and summer months. Notoperlopsis femina Illies, 1963 appeared only in October, while Ceratoperla fazi (Navás 1934) and Pelurgoperla personata Illies, 1963 appeared to be early autumnal species. The most abundant and ubiquitous Ephemeroptera species were Meridialaris chiloeensis, M. diguillina (Demoulin 1955c) and Andesiops torrens (Lugo-Ortiz & McCafferty 1999), whereas Antarctoperla michaelseni (Klapalek 1904), Aubertoperla illiesi (Froehlich 1960), Limnoperla jaffueli (Navás 1928) and Klapopteryx kuscheli were the most frequently collected Plecoptera species. These qualitative abundances and generalized distributions for both mayflies and stoneflies are in agreement with previous reports in the area (Miserendino & Pizzolón 2000). The records of Camelobaetidius, Americabaetis and Traverella were unexpected. Their southern distributional limit was the South American transition zone and the Neotropical region, in the Argentinean provinces of Córdoba and Tucumán, about 1000 km north of our records. These records, together with that of Leptohy- phes Eaton, 1882 (Ephemeroptera), in Somuncurá Plateau, Río Negro province (Muzón & Pessacq 2001) sug- gest a disjunct distributional pattern with interesting biogeographical issues. However, these patterns must be corroborated with extensive sampling in intermediate areas.

TABLE 4. Months and seasons of collection for those species where adults were observed. *Indicates a low quantity of specimens collected at only one sampling site and date. (x) Indicates only one or two specimens collected.

Season Winter Spring Summer Autumn Month Jul. Ag. Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Ephemeroptera Penaphlebia chilensis x Meridialaris chiloeensis x x x x Nousia delicata x x x x x Andesiops ardua x Archethraulodes sp. * x Meridialaris laminata x Meridialaris diguillina x x Metamonius anceps * x Siphlonella sp. x Plecoptera Austronemoura chilena x Antarctoperla michaelseni x x x x x x x Aubertoperla illiesi (x) x x x x x x x Neonemura barrosi (x) (x) x Notoperlopsis femina x Klapopteryx kuscheli x x x x Limnoperla jaffueli x x x x x x Senzilloides panguipulli x x Potamoperla testacea x x Neofulla biloba * x Pelurgoperla personata x Penturoperla barbata * x Rhithroperla rossi * x Udamocercia frantzi x x Austronemoura eudoxiae * x Ceratoperla fazi x x

Some taxa were rare or occasionally recorded, as was the case for Austronemoura eudoxiae, Neofulla biloba (Aubert 1960), Neonemura illiesi, Potamoperla testacea, Alfonsoperla flinti (Plecoptera) and Arche- thraulodes spatulus Pescador & Peters, 1982 (Ephemeroptera). These taxa were represented by only a few specimens at one sampling site. The gripopterygid Potamoperla myrmidon (Mabille 1891) was the only species that appeared through all sampling sites in Chubut River (Miserendino 2006). Similarly, the austroperlid Klapopteryx kuscheli was found in all the sampling sites in Santa Cruz River (province of Santa Cruz) reach- ing the estuarine area on the east of the province (Table 1). This is a curious record, given that this species was always found in the Subantarctic and west limit of the Patagonian subregions, very close to the Andes (Illies 1960a, McLellan 2001). In Patagonia, as well as in many other regions of Argentina and the world over, mayflies and stoneflies are used as indicators of water quality and in biotic indexes (Miserendino & Pizzolón 1999, Pizzolón & Miseren- dino 2000). An adequate knowledge of the species distribution and information on their biotopes improves the

performance of biotic indexes. However, information about systematics, biology and ecology of these taxa in most of Patagonia is scarce. In this sense, collaboration between ecologists and systematists improves the knowledge of these taxa for use in water quality assessment and conservation of these species in Argentina. Even though the present contribution is the result of a limited sampling effort, it increases by 100% the number of Ephemeroptera genera and by 65% the number of Plecoptera species recorded for Chubut province. In addition, it provides nine new country records for Argentina. These results further demonstrate the poor knowledge of the aquatic insect fauna in Patagonia and Chubut province. They also demonstrate the necessity of new and systematic entomological surveys in the area. Patagonian aquatic environments are endangered by the construction of hydroelectric dams, seeding of exotic fished and by poor forestry practices. Long-term poor land use practices have resulted in severe desertification problems in broad areas in the steppe. All these activities threaten the biodiversity of these once pristine and fragile aquatic ecosystems (Pas- cual et al. 1998).

Acknowledgements

Our appreciation is extended to the Administración de Parques Nacionales and to Lic. Claudio Chehébar, Delegación Regional Patagonia, APN, for allowing the specimen collection within Los Alerces National Park, and Dr. Edward DeWalt and the anonymous reviewers for providing several suggestions that greatly improved the manuscript. This work was partially supported by the Argentinean Council of Scientific Research, grant number CONICET, PIP 5733. Special thanks to Dr. Miguel Archangelsky for helpful comments of the English version of the paper.

References

Aubert, J. (1960) Contribution à l´étude des Notomemourinae (Plecoptera) de l´Amériqué du Sud. Bulletin de la Société Entomologique Suisse, 33, 47–64. Claassen, P.W. (1936) New names for stoneflies (Plecoptera). Annals of the Entomological Society of America, 29, 622– 623. Coronato, F.R. & Del Valle, H.F. (1988) Caracterización hídrica de las cuencas hidrográficas de la provincia del Chubut. Publicación Técnica. Cenpat-Conicet. Puerto Madryn. Chubut. Argentina, 183 pp. Demoulin, G. (1955a) Éphéméroptères nouveaux ou rares du Chili. Bulletin de l' Institut Royal des Sciences Naturelles de Belgique, 31, 1–15. Demoulin, G. (1955b) Éphéméroptères nouveaux ou rares du Chili. II. Bulletin de l' Institut Royal des Sciences Naturel- les de Belgique, 31, 1–16. Demoulin, G. (1955c) Éphéméroptères nouveaux ou rares du Chili. III. Bulletin de l' Institut Royal des Sciences Naturel- les de Belgique, 31, 1–30. Demoulin, G. (1966) Contribution à l' étudedes éphéméroptères du Surinam. Bulletin de l' Institut Royal des Sciences Naturelles de Belgique, 42, 1–22. Domínguez, E. (1998) Ephemeroptera. In: Morrone, J. J. y Coscarón, S. (Eds.), Biodiversidad de Artrópodos Argentinos. Una perspectiva biotaxonómica. Ediciones Sur, La Plata, pp. 14–25. Domínguez, E., Hubbard M.D. & Pescador M.L. (1994) Los Ephemeroptera en Argentina. In: Castellanos, Z.A. (Ed.), Fauna de Agua Dulce de la República Argentina. FECIC, Buenos Aires, 33 (1), pp. 1–142. Domínguez, E., Hubbard, M.D., Pescador, M.L. & Molineri, C. (2001) Ephemeroptera. In: H.R. Fernández, E. Domínguez (Eds.), Guía para la Determinación de los Artrópodos Bentónicos Sudamericanos. EudeT, Tucumán, pp.17–54. Eaton, A.E. (1882) An announcement of new genera of Ephemeridae. Entomologist’s Monthly Magazine, 18, 207–208. Eaton, A.E. (1883–1888) A revisional monograhpy of Recent Ephemeridae or Mayflies. Transactions of the Linnean Society of London, Zoology (2) 3, 1–352, pl. 1–65. Edmunds, G.F. Jr. (1948) A new genus of mayflies from western North America (Leptophlebiinae). Proceedings of the

Entomological Society of Washington, 61, 141–146. Enderlein, G. (1909) Klassifikation der Plecopteren sowie Diagnosen neuer Gattungen und Arten. Zoologischer Anzeiger, 34, 385–419. Froehlich, C. G. (1960) Some gripopterygids and notonemurines (Plecoptera) from South America. Lunds Universitets Arsskrift N.F., 56(13), 1–24. Gordon, N.D., Mc Mahon, T.A. & Finlayson, B.L. (1994) Stream Hydrology, an Introduction for Ecologists. Wiley & Sons, New York, 526 pp. Hauer, F.R. & Resh, V.H. 1996. Benthic macroinvertebrates. In: Hauer, F.R. & G.A. Lamberti (Eds), Methods in Stream Ecology. Academic Press, San Diego, California, pp. 339–369 Hynes, H.B.N & Hynes, M.E. (1975) The life history of many of the stoneflies (Plecoptera) of south-eastern mainland Australia. Australian Journal of Marine and Freshwater Research, 27, 61–70. Hollmann, M.E.T. & Miserendino, M.L (2006) Habitat type and life history of the mayflies Metamonius anceps Eaton (Nesameletidae) and Meridialaris chiloeensis Demoulin (Leptophlebidae) in a Patagonian mountain stream. Annales des Limnologie, 42(4), 233–240. Hollmann, M.E.T. & Miserendino M.L. (2008) Life history and emergence patterns of stonefly species in mountain streams of the Futaleufú basin, Patagonia (Argentina). Annales des Limnologie, 44(2): 1-10. Illies, J. (1958) Abranchioperlidae, eine neue Plecopterenfamilie. Investigaciones Zoologicas Chilenas, 164, 26–41. Illies, J. (1960a) Penturoperlidae, eine neue Plecopterenfamilie. Zoologischer Anzeiger, 164, 26–41. Illies, J. (1960b) Archiperlaria, eine neue Unterordnung der Plecopteren. Beiträge zur Entomologie, 10, 661–697. Illies, J. (1961) Südamerikanische Notonemourinae und die Stellung der Unterfamilie im System der Plecopteren. Mittei- lungen der Schweizerischen Entomologischen Gesellschaft, 34, 97–126. Illies, J. (1963) Revision der südamerikanische Gripopterygidae (Plecoptera). Mitteilungen der Schweizerischen Ento- mologischen Gesellschaft, 36, 145–248. Illies, J. (1964) Südamerikanische Perlidae (Plecoptera), besonders au Chile und Argentinien. Beiträge zur Neotropis- chen Fauna, 3, 207–233. Kluge, N. (1992) Cuban mayflies of the family Baetidae (Ephemeroptera). 2. Subgenera Caribaetis subgen. n. and Amer- icabaetis subgen. n. of the genus Baetis s. l. Zoologicheskij Zhurnal, 4, 13–20. Klapálek, F. (1904) Plecopteren. In: Hamburger Magalhaensische Sammelreise, pp 1–30. Léon, R., Bran, D., Collantes, M., Paruelo, J.M. & Soriano, A. (1998) Grandes unidades de vegetación de la Patagonia extra andina. Ecología Austral, 8, 125–144. Lestage, J.A. (1931) Contribution à l’é des Éphéméroptères. VIII. Les Éphéméroptères du Chili. Bulletin et Annales de la Société Entomologique de Belgique, 71, 41–60. Lugo-Ortiz, C.R. & McCafferty W.P. (1996) Taxonomy of the Neotropical genus Americabaetis, new status (Insecta: Ephemeroptera: Baetidae). Studies on Neotropical Fauna and Environment, 31, 156–169. Lugo-Ortiz, C.R. & McCafferty W.P. (1999) Three new genera of small minnow mayflies (Insecta: Ephemeroptera: Baetidae) from the Andes and Patagonia. Studies on Neotropical Fauna and Environment, 34, 88–104. Mabille, J. (1891) Sectio Plecoptera. In: Néuroptéres. Mission scientifique du Cap Horn, 1882–1883, pp. 6–9. Mercado, M. & Elliot, S. (2004) Taxonomic revision of the genus Metamonius Eaton (Nesameletidae: Ephemeroptera), with notes on its biology and distribution. Studies on Neotropical Fauna and Environment, 39(2), 149–157. McLellan, I.D. (2001) A revision of the South American Austroperlidae (Plecoptera). Aquatic Insects, 23(3), 233–251. McLellan, I.D., Miserendino, M.L. & Hollmann, M.E.T. (2006) Two new species of Notoperla (Plecoptera: Gripoptery- gidae) and a redescription of Notoperlopsis femina Illies. Zootaxa, 1140, 63–68. McLellan, I.D., Wais I. & L.I. De Cabo (1990) The first record of stoneflies from the Malvinas/Falkland islands. Aquatic Insetcts, 12(3), 177–180. McLellan, I.D. & Zwick, P. (2007) New species of and keys to South American Gripopterygidae (Plecoptera). Illiesia, 3(4), 20–42. Miserendino, M.L. (1996) Las ninfas de plecóptera de ambientes lóticos del noroeste de Chubut. Naturalia Patagónica. Serie Ciencias Biológicas, 4(1–2), 75–83. Miserendino, M.L. (2001) Macroinvertebrate assemblages in Andean Patagonian rivers and streams. Hydrobiologia, 444,147–158. Miserendino, M.L. (2006) Seasonal and spatial distribution of stoneflies in the Chubut river (Patagonia, Argentina). Hydrobiologia, 568, 263–274. Miserendino, M.L. & Hollmann, M.E.T. (2005) Patrones de emergencia de especies de Plecoptera en ríos del noroeste de Chubut (Patagonia-Argentina). VI Congreso Argentino de Entomología. 12 Septiembre. San Miguel de Tucumán. pp. 191. Miserendino, M.L. & Pizzolón, L.A. (1999) Rapid assessment of river water quality using macroinvertebrates: a family level biotic index for the Patagonic Andean zone. Acta Limnologica Brasiliensia, 11(2), 137–148. Miserendino, M.L. & Pizzolón, L.A. (2000) Macroinvertebrates of a fluvial system in Patagonia: altitudinal zonation and functional structure. Archiv fur Hydrobiologie, 150(1), 55–83.

Miserendino, M.L. & Pizzolón, L.A. (2004) Interactive effects of basin features and land use change on macroinvertebrate communities of headwater streams in the Patagonian Andes. River Research & Applications, 20 (8), 967–982. Morrone, J.J. (2006) Biogeographic areas and transition zones of Latin America and the Caribbean Islands based on pan- biogeographic and cladistic analyses of the entomofauna. Annual Reviews of Entomology, 51, 467–494. Muzón, J. & Bachman, A.O. (1998) Plecoptera. In: Morrone, J. J. y Coscarón, S. (Eds.), Biodiversidad de Artrópodos Argentinos. Una perspectiva biotaxonómica. Ediciones Sur, La Plata, pp. 26–31. Muzón, J. & Pessacq, P. (2001) Nuevos registros de Ephemeroptera para la Meseta de Somuncurá, Río Negro. Revista de la Sociedad Entomológica Argentina, 60 (1–4), 67–78. Navás, L. (1918a) Insectos chilenos. Boletín de la Sociedad Aragonesa de Ciencias Naturales, 17, 212–230. Navás, L. (1918b). Algunos Insectos Neurópteros de la Argentina. Phycis, 4, 81–88. Navás, L. (1919) Insecta Nova. In: Memoires della Pontificia Academia Romana Nuovi Lincei, 2(5), 2–7 Navás, L. (1928) Insectos neotrópicos. Revista Chilena de Historia Natural, 31, 231–232. Navás, L. (1929) Algunos Insectos de Chile. Revista Chilena de Historia Natural, 32, 327–329. Navás, L. (1930) Algunos Insectos de Chile. 3rd. Series. Revista Chilena de Historia Natural, 33, 326–334. Navás, L. (1933) Algunos Insectos de Chile. Revista Chilena de Historia Natural, 37, 231–232. Navás, L. (1934) Insectos suramericanos. Revista de la Academia de Ciencias de Madrid, 31, 155–184. Needham, J.G. & Murphy, H.E. (1924) Neotropical Mayflies. Bulletin of the Loyd Library of Botany, Pharmacy and Materia Medica. 24. Entomological Series, 4, 1–79. Nelson, C.H. (1973) Synopsis of the genus Chilenoperla. The Pan Pacific Entomologist, 49, 315–324. Nieto, C. (2004) South American Baetidae (Ephemeroptera): a New Generic Synonymy. Studies on Neotropical Fauna and Environment, 39(2), 95–101. Pascual, M.A., Orensanz, J.M., Parma, A.M. & Saba, S. (1998) The Patagonian Challenge: Melding Conservation with Development. Conservation Biology: Theory and Practice. In: Fiedler and Kareiva (Eds.), Conservation Biology for the Coming Decade, 2nd edition. Chapman and Hall, London, pp. 410–425. Pescador, M.L. & Peters, W.L. (1980) Two new genera of cool–adapted Leptophlebiidae (Ephemeroptera) from southern South America. Annals of the Entomological Society of Americas, 73, 332–338. Pescador, M.L. & Peters, W.L. (1982) Four new genera of Leptophlebiidae (Ephemeroptera: Atalophlebiinae). Journal of the Kansas Entomological Society, 58, 91–123. Pescador, M.L. & Peters, W.L. (1985) Biosystematics of the genus Nousia from Southern South America (Ephemeroptera: Leptophlebiidae: Atalophlebiinae). Aquatic Insects, 4, 1–19. Pescador, M.L. & Peters, W.L. (1991) Biosystematics of the genus Penaphlebia (Ephemeroptera: Leptophlebiidae: Atalophlebiinae) from South America. Transactions of the American Entomological Society, 117, 1–38. Pictet, F.J. (1841) Historie Naturelle Générale et Particuliere des Insects Névroptères. Famille des Perlides. Genéve, 2 Bände. Pizzolón, L. & Miserendino, M.L. (2000) The performance of two regional biotic indices for running water quality in Northern Patagonian Andes. Acta Limnologica Brasiliensia, 13(1), 11–27. Richardson, J.S. (2001) Life cycle phenology of common detritivores from a temperate rainforest stream. Hydrobiologia, 455, 87–95. Scrimgeour, G.J. (1991) Life history and production of Delatidium (Ephemeroptera: Leptophlebiidae) in an unstable New Zealand river. New Zealand Journal of Marine and Freshwater Research, 25, 93–99. Traver, J.R. & Edmunds, G.F. Jr. (1968) A revision of the Baetidae with spatulate-clawed nymphs (Ephemeroptera). Pacific Insects, 10, 629–677. Ulmer, G. (1920) Neue Ephemeropteren. Archiv für Naturgeschichte, 85A (11), 1–80. Velásquez, S.M. & Miserendino, M.L. (2003) Habitat type and macroinvertebrate assemblages in low order Patagonian streams. Archiv für Hydrobiologie, 158(4), 461–483. Vera, A. (2006) Una nueva especie de Gripopterygidae de Chile, Potamoperla testacea n. sp. (Insecta: Plecoptera). Gay- ana, 70(2), 168–175. Wais, I.R., Villa, A.R. & Cabo, L.I. (1987) Las ninfas de plecópteros de los parques nacionales Nahuel Huapi y Lanin, Argentina. Revista del Museo Argentino de Ciencias Naturales, Hidrobiología, 6(3), 15–21. Zwick, P. (1972) On Neonemura illiesi nov. spec. and some other stoneflies from Chile (Ins. Plecoptera). Studies on Neo- tropical Fauna, 7, 95–100.